The long-distance, unpredictable movement patterns of nomadic species make them challenging to monitor and conserve. Critically endangered regent honeyeaters Anthochaera phrygia once roamed south-eastern Australia in ‘immense flocks’ but now number fewer than 300 wild birds over a vast 300,000 km2 range. Regent honeyeaters are a rare example where extensive monitoring data are now available for a nomadic species, enabling evaluation of the impact of management actions using population viability analysis (PVA). We combined demographic estimates from wild population monitoring in the 1990s, a zoo-based supplementation program and a contemporary range-wide monitoring program to simulate the wild population trajectory under various management and climatic scenarios. Without intervention, our models predicted extinction within 20 years, and showed that management strategies at their current intensity have limited efficacy to prevent extinction. Conservation actions should aim to increase the size and density of the wild population so that Allee effects no longer suppress population growth. Protection of wild regent honeyeater nests is essential as breeding success has declined over recent decades and droughts increasingly reduce breeding opportunities. Our models emphasise the need for zoo-based breeding to bolster the wild population, but show that release of zoo bred birds into the wild only slows the rate of population decline. To recover the wild regent honeyeater population, the next five years will be critical for implementing the most effective conservation strategy. This requires a combination of nest protection and release of zoo-bred birds with improved fitness, predator suppression, habitat protection and increased rates of habitat restoration.

Population viability analysis (PVA) has been used for three decades to assess threats and evaluate conservation options for wildlife populations. What has been learned from PVA on in situ populations are valuable lessons also for assessing and managing viability and sustainability of ex situ populations. The dynamics of individual populations are unpredictable, due to limited knowledge about important factors, variability in the environment, and the probabilistic nature of demographic events. PVA considers such uncertainty within simulations that generate the distribution of likely fates for a population; management of ex situ populations should also take into consideration the uncertainty in our data and in the trajectories of populations. The processes affecting wildlife populations interact, with feedbacks often leading to amplified threats to viability; projections of ex situ populations should include such feedbacks to allow for management that foresees and responds to the cumulative and synergistic threats. PVA is useful for evaluating conservation options only if the goals for each population and measures of success are first clearly identified; similarly, for ex situ populations to contribute maximally to species conservation, the purposes for the population and definitions of sustainability in terms of acceptable risk must be documented. PVA requires a lot of data, knowledge of many processes affecting the populations, modeling expertize, and understanding of management goals and constraints. Therefore, to be useful in guiding conservation it must be a collaborative, trans-disciplinary, and social process. PVA can help integrate management of in situ and ex situ populations within comprehensive species conservation plans.

Breeding programs to conserve diversity are predicated on the assumption that genetic variation in adaptively important traits will be lost in parallel to the loss of variation at neutral loci. To test this assumption, we monitored quantitative traits across 18 generations of Peromyscus leucopus mice propagated with protocols that mirror breeding programs for threatened species. Ears, hind feet, and tails became shorter, but changes were reversible by outcrossing and therefore were due to accumulated inbreeding. Heritability of ear length decreased, because of an increase in phenotypic variance rather than the expected decrease in additive genetic variance. Additive genetic variance in hind foot length increased. This trait initially had low heritability but large dominance or common environmental variance contributing to resemblance among full-sibs. The increase in the additive component indicates that there was conversion of interaction variances to additive variance. For no trait did additive genetic variation decrease significantly across generations. These findings indicate that the restructuring of genetic variance that occurs with genetic drift and novel selection in captivity can prevent or delay the loss of phenotypic and heritable variation, providing variation on which selection can act to adapt populations to captivity and perhaps later to readapt to more natural habitats after release. Therefore, the importance of minimizing loss of gene diversity from conservation breeding programs for threatened wildlife species might lie in preventing immediate reduction in individual fitness due to inbreeding and protecting allelic diversity for long-term evolutionary change, more so than in protecting variation in quantitative traits for rapid re-adaptation to wild environments.

Thousands of small populations are at increased risk of extinction because genetics and evolutionary biology are not well‐integrated into conservation planning–a major lost opportunity for effective actions. We propose that if the risk of outbreeding depression is low, the default should be to evaluate restoration of gene flow to small inbred populations of diploid outbreeding organisms that were isolated by human activities within the last 500 years, rather than inaction. We outline the elements of a scientific‐based genetic management policy for fragmented populations of plants and animals, and discuss the reasons why the current default policy is, inappropriately, inaction.

The risks of inbreeding and outbreeding depression, and the prospects for genetic rescue are often altered in species with different mating systems or modes of inheritance (compared to outbreeding diploids), such as self-incompatible, selffertilizing, mixed mating, non-diploids, and asexual.

Understanding cumulative effects of multiple threats is key to guiding effective management to conserve endangered species. The critically endangered, Southern Resident killer whale population of the northeastern Pacific Ocean provides a data-rich case to explore anthropogenic threats on population viability. Primary threats include: limitation of preferred prey, Chinook salmon; anthropogenic noise and disturbance, which reduce foraging efficiency; and high levels of stored contaminants, including PCBs. We constructed a population viability analysis to explore possible demographic trajectories and the relative importance of anthropogenic stressors. The population is fragile, with no growth projected under current conditions, and decline expected if new or increased threats are imposed. Improvements in fecundity and calf survival are needed to reach a conservation objective of 2.3% annual population growth. Prey limitation is the most important factor affecting population growth. However, to meet recovery targets through prey management alone, Chinook abundance would have to be sustained near the highest levels since the 1970s. The most optimistic mitigation of noise and contaminants would make the difference between a declining and increasing population, but would be insufficient to reach recovery targets. Reducing acoustic disturbance by 50% combined with increasing Chinook by 15% would allow the population to reach 2.3% growth.

The number and geographic location of genetically differentiated populationsmust be identified to determine if fragmented populations require genetic management. Clustering of related genotypes to geographic locations (landscape genetic analyses) is used to determine the number of populations and their boundaries, with the simplest analyses relying on random mating within, but not across populations. Evidence of genetic differentiation among populations indicates either that they have drifted apart (and are likely inbred), or that the populations are adaptively differentiated. The current response when populations are genetically differentiated is usually to recommend separate management, but this is often ill- advised. A paradigm shift is needed where evidence of genetic differentiation among populations is followed by an assessment of whether populations are suffering genetic erosion, whether there are other populations towhich they couldbe crossed, andwhether the crosseswouldbebeneficial or harmful.

Infectious diseases can exert a strong influence on the dynamics of host populations, but it remains unclear why such disease-mediated control only occurs under particular environmental conditions. We used 16 years of detailed field data on invasive European rabbits (Oryctolagus cuniculus) in Australia, linked to individual-based stochastic models and Bayesian approximations, to test whether (i) mortality associated with rabbit haemorrhagic disease (RHD) is driven primarily by seasonal matches/mismatches between demographic rates and epidemiological dynamics and (ii) delayed infection (arising from insusceptibility and maternal antibodies in juveniles) are important factors in determining disease severity and local population persistence of rabbits. We found that both the timing of reproduction and exposure to viruses drove recurrent seasonal epidemics of RHD. Protection conferred by insusceptibility and maternal antibodies controlled seasonal disease outbreaks by delaying infection; this could have also allowed escape from disease. The persistence of local populations was a stochastic outcome of recovery rates from both RHD and myxomatosis. If susceptibility to RHD is delayed, myxomatosis will have a pronounced effect on population extirpation when the two viruses coexist. This has important implications for wildlife management, because it is likely that such seasonal interplay and disease dynamics has a strong effect on long-term population viability for many species.

Summary Species interactions have been largely ignored in extinction risk assessment. However, the black‐footed ferret Mustela nigripes exemplifies a class of endangered species for which strong species interactions cannot be ignored. This species is an obligate predator of prairie dogs Cynomys spp., and sylvatic plague Yersinia pestis epizootics threaten to undermine recovery efforts by functionally eliminating the prey base. Multispecies ‘metamodelling’ techniques offer new opportunities for exploring population dynamics under strong species interdependencies and disease. To investigate ferret extinction risk in plague‐affected landscapes, we simultaneously modelled plague epidemiological processes, prairie dog metapopulation dynamics and ferret demographic responses. Ferret population dynamics were investigated at an important release site (Conata Basin in South Dakota) and for 500 artificial prey landscapes spanning a wide range of realistic colony configurations (e.g. total area, # colonies, spatial clustering) and demographic characteristics. Our simulation models indicate that ferrets are unlikely to persist through episodes of plague at Conata Basin unless they can access prey resources from a wider region or unless management actions can otherwise substantially reduce plague transmission. We show that large, diffuse prairie dog metapopulations (those with colonies spread over a region >2500 km2) are most likely to support ferret populations in plague‐affected landscapes. Our results also highlight the potential importance of metapopulation connectivity in fuelling plague epizootics and thereby imperilling black‐footed ferret conservation efforts. We describe a cycle (c. 5‐ to 25‐year period) of plague‐driven population crashes that is an emergent property of our models, and which can destabilize ferret populations. Synthesis and applications. On the basis of our models, we conclude that few North American prairie dog complexes cover sufficient land area to sustain black‐footed ferret populations through plague‐driven crashes in prey abundance. Consequently, our results underscore the importance of working with many constituents to conserve large prairie dog landscapes in addition to continued development of plague mitigation tools. In addition, the strong relationship between plague‐induced oscillatory prey cycles and predator population persistence highlights the potential conservation benefits of imposing strategic barriers to connectivity in areas over which plague outbreak cycles are strongly synchronous.

Genetic relatedness between individuals is an important measure in many areas of biology. However, some relatedness measures for use with molecular (allele) data assume that the individuals themselves are not inbred. Here, we present a new measure of relatedness based on the different modes of identity-by-descent for alleles that has an upper bound of 1 even when the individuals being compared are themselves inbred. This new measure is compared to several other measures of relatedness using several simple examples and pedigree data from the wolf population in Isle Royale National Park.

The goal of captive breeding programmes is often to maintain genetic diversity until re-introductions can occur. However, due in part to changes that occur in captive populations, approximately one-third of re-introductions fail. We evaluated genetic changes in captive populations using microsatellites and mtDNA. We analysed six populations of white-footed mice that were propagated for 20 generations using two replicates of three protocols: random mating (RAN), minimizing mean kinship (MK) and selection for docility (DOC). We found that MK resulted in the slowest loss of microsatellite genetic diversity compared to RAN and DOC. However, the loss of mtDNA haplotypes was not consistent among replicate lines. We compared our empirical data to simulated data and found no evidence of selection. Our results suggest that although the effects of drift may not be fully mitigated, MK reduces the loss of alleles due to inbreeding more effectively than random mating or docility selection. Therefore, MK should be preferred for captive breeding. Furthermore, our simulations show that incorporating microsatellite data into the MK framework reduced the magnitude of drift, which may have applications in long-term or extremely genetically depauperate captive populations.

Measurements of size and asymmetry in morphology might provide early indications of damaging effects of inbreeding or other genetic changes in conservation breeding programs. We examined the effects of inbreeding on size and fluctuating asymmetry (FA) in skull and limb bone measurements in experimental populations of three subspecies of Peromyscus polionotus mice that had previously been shown to suffer significant reductions in reproductive success when inbred. Inbreeding caused significant depression in mean size in two of the subspecies (P. p. rhoadsi and P. p. subgriseus), but the effects were smaller in the third (P. p. leucocephalus). Inbreeding caused an increase in FA of just one of eight bilateral traits in one subspecies (P. p. rhoadsi). Inbreeding depression in size was more easily detected than the effects of inbreeding on FA. FA may be much less sensitive to inbreeding and other stresses than are more direct measures of fitness such as reproductive output and body mass growth rate. Given the large sample sizes and statistical complexity required to assess changes to typically very small levels of FA in captive populations, FA will not likely provide a useful measure of inbreeding depression in captive populations.

Wildlife population models have been criticized for their narrow disciplinary perspective when analyzing complexity in coupled biological - physical - human systems. We describe a "metamodel" approach to species risk assessment when diverse threats act at different spatiotemporal scales, interact in non-linear ways, and are addressed by distinct disciplines. A metamodel links discrete, individual models that depict components of a complex system, governing the flow of information among models and the sequence of simulated events. Each model simulates processes specific to its disciplinary realm while being informed of changes in other metamodel components by accessing common descriptors of the system, populations, and individuals. Interactions among models are revealed as emergent properties of the system. We introduce a new metamodel platform, both to further explain key elements of the metamodel approach and as an example that we hope will facilitate the development of other platforms for implementing metamodels in population biology, species risk assessments, and conservation planning. We present two examples - one exploring the interactions of dispersal in metapopulations and the spread of infectious disease, the other examining predator-prey dynamics - to illustrate how metamodels can reveal complex processes and unexpected patterns when population dynamics are linked to additional extrinsic factors. Metamodels provide a flexible, extensible method for expanding population viability analyses beyond models of isolated population demographics into more complete representations of the external and intrinsic threats that must be understood and managed for species conservation.

For the last 30 years, cooperative management of irreplaceable animal populations in zoos and aquariums has focused primarily on the goal of minimizing genetic decay within defined time frames, and large advances have been made in technologies to optimize genetic management of closed populations. However, recent analyses have shown that most zoo programs are not projected to meet their stated goals. This has been described as a lack of achieving "sustainability" of the populations, yet by definition a goal of managed decay is not a plan for sustainability. True sustainability requires management of the resource in manner that does not deplete its value for the future. Achieving such sustainability for many managed populations may require changing from managing isolated populations to managing populations that are part of a broader metapopulation, with carefully considered exchange between populations across a spectrum of ex situ to in situ. Managing zoo populations as components of comprehensive conservation strategies for the species will require research on determinants of various kinds of genetic, physiological, behavioral, and morphological variation and their roles in population viability, development of an array of management techniques and tools, training of population managers in metapopulation management and integrated conservation planning, and projections of impacts of management strategies on the viability of the captive populations and all populations that are interactively managed or affected. Such a shift in goals and methods would result in zoo population management being an ongoing part of species conservation rather than short-term or isolated from species conservation. Zoo Biol. 32:19-26, 2013. © 2012 Wiley Periodicals, Inc.

Restoring connectivity between fragmented populations is an important tool for alleviating genetic threats to endangered species. Yet recovery plans typically lack quantitative criteria for ensuring such population connectivity. We demonstrate how models that integrate habitat, genetic, and demographic data can be used to develop connectivity criteria for the endangered Mexican wolf (Canis lupus baileyi), which is currently being restored to the wild from a captive population descended from 7 founders. We used population viability analysis that incorporated pedigree data to evaluate the relation between connectivity and persistence for a restored Mexican wolf metapopulation of 3 populations of equal size. Decreasing dispersal rates greatly increased extinction risk for small populations (<150-200), especially as dispersal rates dropped below 0.5 genetically effective migrants per generation. We compared observed migration rates in the Northern Rocky Mountains (NRM) wolf metapopulation to 2 habitat-based effective distance metrics, least-cost and resistance distance. We then used effective distance between potential primary core populations in a restored Mexican wolf metapopulation to evaluate potential dispersal rates. Although potential connectivity was lower in the Mexican wolf versus the NRM wolf metapopulation, a connectivity rate of >0.5 genetically effective migrants per generation may be achievable via natural dispersal under current landscape conditions. When sufficient data are available, these methods allow planners to move beyond general aspirational connectivity goals or rules of thumb to develop objective and measurable connectivity criteria that more effectively support species recovery. The shift from simple connectivity rules of thumb to species-specific analyses parallels the previous shift from general minimum-viable-population thresholds to detailed viability modeling in endangered species recovery planning.

Many wildlife species are propagated in captivity as models for behavioral, physiological, and genetic research or to provide assurance populations to protect threatened species. However, very little is known about how animals evolve in the novel environment of captivity. The histories of most laboratory strains are poorly documented, and protected populations of wildlife species are usually too small and too short-term to allow robust statistical analysis. To document the evolutionary change in captive breeding programs, we monitored reproduction and behavior across 18 generations in six experimental populations of Peromyscus leucopus mice started from a common set of 20 wild-caught founders. The mice were propagated under three breeding protocols: a strategy to retain maximal genetic diversity, artificial selection against stereotypic behaviors that were hypothesized to reflect poor adaptation to captivity, and random bred controls. Two replicates were maintained with each protocol, and inter-replicate crosses at generations 19 and 20 were used to reverse accumulated inbreeding. We found that one of the stereotypic behaviors (repetitive flipping) was positively associated with reproductive fitness, while the other (gnawing) was relatively invariant. Selection to reduce these stereotypic behaviors caused marked reduction in reproduction, and populations not under artificial selection to reduce these behaviors responded with large increases in flipping. In non-selected populations, there was rapid evolution toward much higher proportion of pairs breeding and more rapid conception. Litter size, pup survival, and weaning mass all declined slowly, to the extent that would be predicted based on inbreeding depression. Inter-crossing between replicate populations reversed these declines in fitness components but did not reverse the changes in behavior or the accelerated breeding. These findings indicate that adaptation to captivity can be rapid, affecting reproductive patterns and behaviors, even under breeding protocols designed to minimize the rate of genetic change due to random drift and inadvertent selection.

The 26 definitions of species often yield different numbers of species and disparate groupings, with financial, legal, biological and conservation implications. Using conservation genetic considerations, we demonstrate that different species concepts have a critical bearing on our ability to conserve species. Many species of animals and plants persist as small isolated populations suffering inbreeding depression, loss of genetic diversity, and elevated extinction risks. Such small populations usually can be rescued by restoring gene flow, but substantial genetic drift effects can lead them to be classified as distinct species under the diagnostic phylogenetic species concept. Minimum harm to fitness is done and maximum potential fitness and evolutionary potential benefits accrue when reproductive isolation (pre- and/or post-zygotic) is used as the criterion to define distinct species. For sympatric populations, distinct species are diagnosed by very limited gene flow. For allopatric populations, both minimal gene flow and evidence of reduced reproductive fitness in crosses (or effects predicted from adaptive differentiation among populations and/or fixed chromosomal differences) are required to satisfy conservation issues. Species delineations based upon the biological and differential fitness species concepts meet the above requirements. Conversely, if species are delineated using the diagnostic phylogenetic species concept, genetic rescue of small genetically isolated populations may require crosses between species, with consequent legal and regulatory ramifications that could preclude actions to prevent extinction. Consequently, we conclude that the diagnostic phylogenetic species concept is unsuitable for use in conservation contexts, especially for classifying allopatric populations. (C) 2012 Elsevier Ltd. All rights reserved.